Spatial point pattern analysis of available and exploited resources

A patchy spatial distribution of resources underpins many models of population regulation and species coexistence, so ecologists require methods to analyse spatially‐explicit data of resource distribution and use. We describe a method for analysing maps of resources and testing hypotheses about species' distributions and selectivity. The method uses point pattern analysis based on the L‐function, the linearised form of Ripley's K‐function. Monte Carlo permutations are used for statistical tests. We estimate the difference between observed and expected values of L(t), an approach with several advantages: 1) The results are easy to interpret ecologically. 2) It obviates the need for edge correction, which has largely precluded the use of L‐functions where plot boundaries are “real”. Including edge corrections may lead to erroneous conclusions if the underlying assumptions are invalid. 3) The null expectation can take many forms, we illustrate two models: complete spatial randomness (to describe the spatial pattern of resources in the landscape) and the underlying pattern of resource patches in the landscape (akin to a neutral landscape approach). The second null is particularly useful to test whether spatial patterns of exploited resource points simply reflect the spatial patterns of all resource points. We tested this method using over 100 simulated point patterns encompassing a range of patterns that might occur in ecological systems, and some very extreme patterns. The approach is generally robust, but Type II decision errors might arise where spatial patterns are weak and when trying to detect a clumped pattern of exploited points against a clumped pattern of all points. An empirical example of an intertidal lichen growing on barnacle shells illustrates how this technique might be used to test hypotheses about dispersal mechanisms. This approach can increase the value of survey data, by permitting quantification of natural resource patch distribution in the landscape as well as patterns of resource use by species.     

A new non‐parametric method for analyzing replicated point patterns in ecology

Most ecological studies that involve point pattern analyses are based on a single plot, which prevent the separation of the effects of various processes that could act simultaneously, as well as limiting the conclusions that can be extracted from these studies. However, considering the spatial distribution of individuals in several plots as replicates of the same process could help to differentiate its specific effects from those of other confounding processes. Thus, we introduce a new method for analyzing spatial point patterns that are replicated according to a two‐factorial design. By summarizing the spatial patterns as K‐functions, the proposed method computes the average K‐functions for each level of the two factors (i.e. predictors) and for each combination of levels, before estimating the sum of squared deviations from the overall mean K‐function. Inferences of the strength of the relationship between the predictors, their interaction, and the spatial structure are made based on a non‐parametric bootstrap procedure, which considers the dependency among spatial scales. We illustrate the proposed approach based on an analysis of the effects of altitude (with two levels: low and high) and slope (with two levels: flat and steep slopes) on the spatial pattern of Croton wagneri, a dominant shrub in an Andean dry scrubland. Our method detected a significant effect of the interaction between slope and altitude, which could not have been detected using current point pattern analysis methodology. The prevalence of single‐plot analysis in ecological studies may be due to a lack of familiarity with appropriate methods for replicated point patterns, as well as the greater complexity of these methods and the absence of appropriate software. Our approach can be applied to a significant number of ecological questions while maintaining a simple, understandable, and easily reportable methodological framework.     

Spatial patterns of an endemic Mediterranean palm recolonizing old fields

Throughout Europe, increased levels of land abandonment lead to (re)colonization of old lands by forests and shrublands. Very little is known about the spatial pattern of plants recolonizing such old fields. We mapped in two 21–22‐ha plots, located in the Doñana National Park (Spain), all adult individuals of the endozoochorous dwarf palm Chamaerops humilis L . and determined their sex and sizes. We used techniques of spatial point pattern analysis (SPPA) to precisely quantify the spatial structure of these C. humilis populations. The objective was to identify potential processes generating the patterns and their likely consequences on palm reproductive success. We used (1) Thomas point process models to describe the clustering of the populations, (2) random labeling to test the sexual spatial segregation, and (3) mark correlation functions to assess spatial structure in plant sizes. Plants in both plots showed two critical scales of clustering, with small clusters of a radius of 2.8–4 m nested within large clusters with 38–44 m radius. Additional to the clustered individuals, 11% and 27% of all C. humilis individuals belonged to a random pattern that was independently superimposed to the clustered pattern. The complex spatial pattern of C. humilis could be explained by the effect of different seed‐dispersers and predators' behavior and their relative abundances. Plant sexes had no spatial segregation. Plant sizes showed a spatial aggregation inside the clusters, with a decreasing correlation with distance. Clustering of C. humilis is strongly reliant on its seed dispersers and stressful environmental conditions. However, it seems that the spatial patterns and dispersal strategies of the dwarf palm make it a successful plant for new habitat colonization. Our results provide new information on the colonization ability of C. humilis and can help to develop management strategies to recover plant populations.     

Bayesian Wombling for Spatial Point Processes

Summary In many applications involving geographically indexed data, interest focuses on identifying regions of rapid change in the spatial surface, or the related problem of the construction or testing of boundaries separating regions with markedly different observed values of the spatial variable. This process is often referred to in the literature as boundary analysis or wombling. Recent developments in hierarchical models for point‐referenced (geostatistical) and areal (lattice) data have led to corresponding statistical wombling methods, but there does not appear to be any literature on the subject in the point‐process case, where the locations themselves are assumed to be random and likelihood evaluation is notoriously difficult. We extend existing point‐level and areal wombling tools to this case, obtaining full posterior inference for multivariate spatial random effects that, when mapped, can help suggest spatial covariates still missing from the model. In the areal case we can also construct wombled maps showing significant boundaries in the fitted intensity surface, while the point‐referenced formulation permits testing the significance of a postulated boundary. In the computationally demanding point‐referenced case, our algorithm combines Monte Carlo approximants to the likelihood with a predictive process step to reduce the dimension of the problem to a manageable size. We apply these techniques to an analysis of colorectal and prostate cancer data from the northern half of Minnesota, where a key substantive concern is possible similarities in their spatial patterns, and whether they are affected by each patient's distance to facilities likely to offer helpful cancer screening options.     

Causes of pattern in plant communities where environmental change is rapid and species longevity is short

Questions : To what extent can spatial structure and its causes be determined in a highly disturbed environment? What are the main determinants of pattern and are these species‐specific? How much do spatial patterns change over generations? Location : Wimmera region of southern Australia. Methods : Broad‐leaved weeds were counted in 225 000 contiguous 20‐cm square quadrats. A substantial number of these quadrats were recorded again after two and four years. An hierarchical ‘adaptive analysis’ approach was used to select spatial analytical methods to examine specific aspects of pattern and variation in pattern from year to year. Results : Patterns varied among species and included both dense and sparse patches surrounded by areas of zero density, diffuse gradations of density and clear anisotropy. Patterns in Erodium botrys and Oxalis pes‐caprae persisted over years, whereas patterns in Arctotheca calendula were less pronounced and varied over time. Edaphic factors appeared to have only a minor influence over the spatial distribution of the weed community as a whole. In Oxalis pes‐caprae, whose patches were hypothesized to have been shaped by cultivation, there was no spread in four years, despite further tillage. Outlying plants of O. pes‐caprae failed to establish new patches, even in the year of greatest population increase. Little evidence of localised recruitment events was found. Conclusions : Despite repeated annual disturbances by natural and anthropogenic mechanisms, clear and interpretable spatial structure develops in annual weeds over a range of spatial resolutions. Adaptive analysis is a useful approach to the characterization of such patterns.     

Natural regeneration of forest related to the spatial structure of trees: A study of two forest communities in Western Carpathians,southern Poland

The existence of a relationship between the spatial pattern of trees and the distribution of young individuals beneath the canopy has been tested in the beech (Fagus sylvatica) and spruce (Picea abies) — fir (Abies alba) forests in the mountainous region, using two different methods. The first method was the analysis of spatial pattern of individuals, the second one was based on calculating sums of influences of all trees occurring within analysed plot on a given point on the forest floor. Results of spatial pattern analyses were surprisingly consistent: almost all mature trees and seedlings didplayed a random pattern of spatial arrangement. However, there is a clear, although statistically insignificant tendency towards uniformity of spatial pattern with increasing sizes of analysed trees. Results of comparing sums of influences on regularly distributed points with sums of influences on seedlings or saplings revealed no tendency in forest regeneration to concentrate in places, where the sums were smaller than the average for a plot. This, coupled with the dominance of random spatial pattern of trees, suggests, that viewed on a small spatial scale, influence of competition among forest trees on their spatial arrangement is obscured by other factors, which are not closely related to the distribution of individuals.     

Structural segregation and scales of spatial dependency in Abies amabilis forests

Question: Is a mosaic structure apparent in the spatial distribution of trees in old‐growth Abies amabilis forests? Location: Montane forests of the western Cascade Range, Washington, USA. Methods: Maps of tree locations were created for study areas located in two, 300‐year old stands and a single 600‐year old stand. Stand structure parameters were calculated using several subsample quadrats sizes (56.25 ‐ 306.25 m²), which were drawn randomly with replacement at a density of 250 quadrats per ha from the stem maps in the computing environment. Spatial cross‐covariance functions between different canopy strata were estimated using the spline cross‐correlogram. Results: Negative spatial correlation (segregation) between subcanopy tree density and areas of high overstorey occupancy was detected. Understorey and midstorey tree densities were positively spatially correlated. These general trends were apparent across the range of observational scales investigated. Significant spatial correlation between canopy strata was observed at spatial scales of 12 ‐ 44 m and extended to the largest scales in the 600‐year old stand. Conclusion: The observed spatial segregation between canopy strata supports the hypothesis that old A. amabilis forests form fine‐scale structural mosaics. Structural segregation at small scales may be due to competitive interactions as well as exogenous forcing of tree locations (e.g. by mortality due to pathogens or disturbance), however segregation at large scales in the 600‐year old stand is likely due to exogenous factors alone. This study reinforces the idea that horizontal heterogeneity is an emergent property of old‐growth forests.     

Beyond the tropics: forest structure in a temperate forest mapped plot

Question: How do the diversity, size structure, and spatial pattern of woody species in a temperate (Mediterranean climate) forest compare to temperate and tropical forests? Location: Mixed evergreen coastal forest in the Santa Cruz Mountains, California, USA. Methods: We mapped, tagged, identified, and measured all woody stems (≥1 cm diameter) in a 6‐ha forest plot, following Center for Tropical Forest Science protocols. We compared patterns to those found in 14 tropical and 12 temperate forest plots. Results: The forest is dominated by Douglas‐fir (Pseudotsuga menziesii) and three species of Fagaceae (Quercus agrifolia, Q. parvula var. shrevei, and Lithocarpus densiflorus), and includes 31 woody species and 8180 individuals. Much of the diversity was in small‐diameter shrubs, treelets, and vines that have not been included in most other temperate forest plots because stems <5‐cm diameter had been excluded from study. The density of woody stems (1363 stems ha^?1) was lower than that in all but one tropical plot. The density of large trees (diameter ≥30 cm) and basal area were higher than in any tropical plot. Stem density and basal area were similar to most other temperate plots, but were less than in low‐diversity conifer forests. Rare species were strongly aggregated, with the degree of aggregation decreasing with abundance so that the most common species were significantly more regular than random. Conclusions: The patterns raise questions about differences in structure and dynamics between tropical and temperate forests; these need to be confirmed with additional temperate zone mapped plots that include small‐diameter individuals.     

The spatial relationship between post‐crop remaining trees and the establishment of saplings in Pinus sylvestris stands in Spain

Question: What is the spatial relationship between remaining trees and the establishment and development of recruited saplings? Location: The Pinus sylvestris forest Pinar de Valsaín, in the Sistema mountain range (central Spain). Methods: Three 0.5 ha plots have been analysed. The saplings were located in a 2 m x 2 m grid, characterizing their spatial pattern through a nested ANO VA. The spatial pattern of stems was analysed using the L(d) function. To analyse the spatial relationship between stems belonging to different cohorts, the intertype L_rs(d) function was used. Finally a new function K_rx(d) is presented as a method to analyse the relationship between the spatial distribution of stems and the sapling density (a sampled continuous variable). Results: The mother trees show cluster pattern at scales of ca. 12 m ‐ 22 m, leading to a spatial pattern at 14 m–16 m for the saplings during the regeneration period. At the beginning of the shelter phase, saplings less than 1.30 m in height show spatial repulsion from the old crop at distances above 10 m, whereas taller saplings show repulsion at shorter distances, due to the suppression of sapling development near the mother trees. At the end of the regeneration period, saplings < 1.30 m appear under the last remaining mother tree canopies. Conclusions: In the stands analysed, located at the southern limit of Pinus sylvestris distribution, this species behaves as half‐shade tolerant. This study shows that the K_rx(d) function might be widely applied to analyse the relationship between patterns that occur at different scales or between a point pattern and a continuous variable, being a useful tool for analysing some forest processes.     

长白山次生杨桦林种群空间点格局及密度制约效应

以长白山5.2 hm²次生杨桦林样地为研究对象,利用空间点格局分析的双相关函数g（r）以及随机标签零模型和案例-对照设计法,探讨了树木种群空间格局及其密度制约效应。研究结果表明：生境异质性对不同生活型以及不同树种的成熟个体在大尺度范围上具有强烈影响。剔除生境异质性带来的影响后,在14个常见树种中有12个树种在小尺度上呈显著的空间聚集分布格局;随着空间尺度的增加聚集性分布树种数量急剧下降,在18 m尺度上聚集率下降到0。全部14个常见树种均呈现出显著的密度制约效应,表明密度制约是调节温带森林树木种群空间结构的主要作用机制。此外,密度制约与物种多度呈负相关,与亚林层和灌木层树种的同种聚集强度呈正相关。并且随着空间尺度增大受密度制约影响的树种百分比逐渐减小,14个常见树种中有11个在0-1 m处达到最大的密度制约强度。     

Within‐field spatial distribution patterns of corn planthopper,Peregrinus maidis,and severity of hopperburn and Maize mosaic virus symptoms as influenced by sunn hemp intercropping

Habitat management (e.g., intercropping) may alter within‐field spatial distribution patterns of herbivores, from a typical pattern as observed in a monoculture, and may influence patterns of crop injury. Field trials were conducted to study the effect of intercropping maize, Zea mays L. (Poaceae), with sunn hemp, Crotalaria juncea L. (Fabaceae) strips on within‐field spatial distribution patterns of corn planthopper, Peregrinus maidis (Ashmead) (Hemiptera: Delphacidae), and combined severity of hopperburn and Maize mosaic virus (MMV) (Rhabdoviridae: Nucleorhabdovirus) symptoms. In each field trial, spatially explicit data on P. maidis counts and ratings of severity of symptoms were obtained by sampling maize plants at weekly intervals. These data were used to examine the spatial patterns of P. maidis and severity of symptoms in maize‐intercropped and monoculture plots with Spatial Analysis for Distance IndicEs (SADIE) methodology. Spatial aggregation patterns of P. maidis in each treatment plot were not consistent among the field trials and tended to be mediated by their population densities. Interpolation of local cluster indices showed that P. maidis were more often aggregated at the field edges, irrespective of treatment. At times of MMV incidence in field trials (fall 2010 and spring 2011), the patch clusters of P. maidis and symptomatic plants were located at the field edges, but were spatially unassociated in both treatment plots. The results provided an approximation of the unpredictability of P. maidis spatial patterns at different population densities and their association with severity of symptoms in two maize‐cropping systems. However, the gap clusters of symptomatic plants were primarily located at the field interiors and were larger in intercropped than in monoculture plots. Such spatial pattern of symptomatic plants resulted in the reduced incidence of MMV in the intercropped plot compared with the monoculture plot, suggesting intercropping sunn hemp can be a useful tool in the management of MMV in maize fields.     

A new digital method of data collection for spatial point pattern analysis in grassland communities

A major objective of plant ecology research is to determine the underlying processes responsible for the observed spatial distribution patterns of plant species. Plants can be approximated as points in space for this purpose, and thus, spatial point pattern analysis has become increasingly popular in ecological research. The basic piece of data for point pattern analysis is a point location of an ecological object in some study region. Therefore, point pattern analysis can only be performed if data can be collected. However, due to the lack of a convenient sampling method, a few previous studies have used point pattern analysis to examine the spatial patterns of grassland species. This is unfortunate because being able to explore point patterns in grassland systems has widespread implications for population dynamics, community‐level patterns, and ecological processes. In this study, we developed a new method to measure individual coordinates of species in grassland communities. This method records plant growing positions via digital picture samples that have been sub‐blocked within a geographical information system (GIS). Here, we tested out the new method by measuring the individual coordinates of Stipa grandis in grazed and ungrazed S. grandis communities in a temperate steppe ecosystem in China. Furthermore, we analyzed the pattern of S. grandis by using the pair correlation function g(r) with both a homogeneous Poisson process and a heterogeneous Poisson process. Our results showed that individuals of S. grandis were overdispersed according to the homogeneous Poisson process at 0–0.16 m in the ungrazed community, while they were clustered at 0.19 m according to the homogeneous and heterogeneous Poisson processes in the grazed community. These results suggest that competitive interactions dominated the ungrazed community, while facilitative interactions dominated the grazed community. In sum, we successfully executed a new sampling method, using digital photography and a geographical information system, to collect experimental data on the spatial point patterns for the populations in this grassland community.     

Dispersal constraints and fine‐scale spatial genetic structure in two earthworm species

The limited dispersal ability of earthworms is expected to result in marked genetic isolation by distance and remarkable spatial patterns of genetic variation. To test this hypothesis, we investigated, using microsatellite loci, the spatial genetic structure of two earthworm species, Allolobophora chlorotica and Aporrectodea icterica, in two plots of less than 1 ha where a total of 282 individuals were collected. We used spatial autocorrelation statistics, partial Mantel tests of isolation‐by‐distance (IBD) and isolation‐by‐resistance (IBR), and Bayesian test of clustering to explore recent patterns involved in the observed genetic structure. For A. icterica, a low signal of genetic structure was detected, which may be explained by an important dispersal capacity and/or by the low polymorphism of the microsatellite loci. For A. chlorotica, a weak, but significant, pattern of IBD associated with positive autocorrelation was observed in one of the plots. In the other plot, which had been recently ploughed, two genetically differentiated clusters were identified. These results suggest a spatial neighbourhood structure in A. chlorotica, with neighbour individuals that tend to be more genetically similar to one another, and also highlight that habitat perturbation as a result of human activities may deeply alter the genetic structure of earthworm species, even at a very small scale. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 335–347.     

Variations in resistance to canopy disturbances and their interactions with the spatial structure of major species in a cool‐temperate forest

Question: How does typhoon‐related disturbance (more specifically, disturbance in the understorey due to tree‐fall and branch‐fall) affect different species mortality rates in a vertically well‐structured forest community? Location: Cool‐temperate, old‐growth forest in the Daisen Forest Reserve, Japan. Methods: We investigated the canopy dynamics and mortality rate trends of trees ≥5 cm diameter at breast height in a 4‐ha study plot, and analysed the effects of tree diameter and spatial structure on the mortality risks for major tree species in the understorey. Results: Significant differences were found in the mortality rates and proportions of injured dead stems between census periods, which were more pronounced in the understorey than in the canopy. Acer micranthum, which showed increased mortality during typhoon disturbance periods, had a clumped distribution. In contrast, Acer japonicum and Viburnum furcatum, which showed similar mortality rates between census periods, had a loosely clumped spatial distribution and a negative association with canopy trees, respectively. In the understorey stems of Acanthopanax sciadophylloides and Fagus crenata, whose spatial distribution patterns depended on canopy gaps, significant increases in mortality rates were observed only during severe typhoon‐related disturbance periods. Conclusions: The sensitivity of trees to typhoon‐related canopy disturbance is more pronounced in the lower layers of vertically structured forest communities. Differences in mortality patterns generated through the combined effects of spatial variation in disturbance regime and species‐specific spatial distribution patterns (spatial aggregation, association with canopy trees, and canopy gap dependency) contribute to the co‐existence of understorey species in forest communities that are subject to typhoon‐related disturbance.     

Woody vegetation spatial patterns in a semi-arid savanna of Burkina Faso,West Africa

Spatial patterns of woody individuals were studied in a semi-arid savanna of West Africa located in Burkina Faso at and around 14° 12 N and 2° 27 W. The study was based upon a 10.24 ha plot within which individuals were mapped. Spatial pattern analysis was carried out using second order characteristics of point processes as K functions and pair correlations. The overall density amounted to 298 individuals ha^-1. The most abundant species were Combretum micranthum G. Don., Grewia bicolor Juss. and Pterocarpus lucens Lepr. Anogeissus leiocarpus (D.C.) G. et Perr. was also an important constituant of this vegetation type, owing to its taller stature. Clumped spatial distributions were identified for all species except for two, for which complete spatial randomness (CSR) was found (including P. lucens, a dominant woody plant). No regular pattern was found even when tall individuals were considered alone. Aggregation dominates interspecific relationships, resulting in multispecific clumps and patches. The overall aggregation pattern was constituted by two different structures. A coarse-grain pattern of ca. 30–40 m was based on edaphic features, and expresses the contrast between sparse stands on petroferric outcrops and denser patches on less shallow soils. A finer-grain pattern made of clumps ca. 5–10 m wide, with no obvious relation to pre-existing soil heterogeneity. There was no overall pattern for saplings (between 0.5 m and 1.5 m in height) irrespective of species, and thus no obvious common facilitation factor. For species with a high recruitment level there was no significant relationship between mature adult and saplings. The only case of clumped saplings with randomly distributed adults was found in P. lucens. However, this cannot be unequivocally interpreted as density dependent regulation since the existence of such a process was not consistent with the spatial distribution of dead P. lucens individuals (victims of the last drought). The mean density around dead P. lucens was lower than around surviving ones, indicating that the last drought tended to reinforce clumping rather than promote a regular pattern of trees. Spatial pattern analysis yielded no evidence supporting a hypothesis of stand density regulation through competition between individuals. Other processes, as surface sealing of bare soils or insufficient recruitment, may play a more important role in preventing a savanna-like vegetation from turning into denser woodlands or thickets.     

The effects of plant dispersion and prey density on parasitism rates in a naturally patchy habitat

Despite extensive research on parasitoid-prey interactions and especially the effects of heterogeneity in parasitism on stability, sources of heterogeneity other than prey density have been little investigated. This research examines parasitism rates by three parasitoid species in relationship to prey density and habitat spatial pattern. The herbivore Itame andersoni (Geometridae) inhabits a subdivided habitat created by patches of its host plant, Dryas drummondii, in the Wrangell Mountains of Alaska. Dryas colonizes glacial moraines and spreads clonally to form distinct patches. Habitat subdivision occurs both on the patch scale and on the larger spatial scale of sites due to patchy successional patterns. Itame is attacked by three parasitoids: an ichneumonid wasp (Campoletis sp.), a braconid wasp (Aleiodes n. sp.), and the tachinid fly (Phyrxe pecosensis). I performed a large survey study at five distinct sites and censused Itame density and parasitism rates in 206 plant patches for 1–3 years. Parasitism rates varied with both plant patch size and isolation and also between sites, and the highest rates of overall parasitism were in the smallest patches. However, the effects of both small- and large-scale heterogeneity on parasitism differed for the three parasitoid species. There was weak evidence that Itame density was positively correlated with parasitism for the braconid and tachinid at the patch scale, but density effects differed for different patch sizes, patch isolations, and sites. At the site scale, there was no evidence of positive, but some indication of negative density-dependent parasitism. These patterns do not appear to be driven by negative interactions between the three parasitoid species, but reflect, rather, individual differences in habitat use and response to prey density. Finally, there was no evidence that parasitism strongly impacted the population dynamics of Itame. These results demonstrate the importance of considering habitat pattern when examining spatial heterogeneity of parasitism and the impacts of parasitoids. Received: 3 June 1999 / Accepted: 4 October 1999     

108 years of change in spatial pattern following selective harvest of a Pinus ponderosa stand in northern Arizona,USA

Questions: How did an initial tree harvest in 1894 influence the spatial and temporal patterns of Pinus ponderosa recruitment? How do these patterns compare to our understanding of P. ponderosa stand dynamics prior to Euro‐American settlement? How might spatial pattern information, particularly with respect to patch characteristics, inform current restoration and management practices? Location: A 2.59‐ha permanent sample plot in the Fort Valley Experimental Forest, Flagstaff, Arizona. The plot was selectively harvested in 1894 and measured in 1909 and 2002. Methods: We used historical stem‐map and ledger data, contemporary data, and dendrochronological techniques to reconstruct stand structure (tree size, age, location) in three scenarios: (1) unharvested (1909), (2) harvested (1909), and (3) contemporary (2002). We used Clark and Evans' R, Ripley's K(t) univariate analysis, and correlogram analysis to assess the spatial pattern in each scenario. We also used Ripley's K₁₂(t) bivariate analysis and tree age data to examine spatial and temporal recruitment patterns as observed in the contemporary scenario. Results and Conclusions: The unharvested stand was aggregated at scales up to 28 m. The selective harvest accentuated the spatial patchiness of the stand in 1909 and changed spatial patterns by homogenizing tree size within patches. By 2002, the stand was a single patch dominated by small trees. Post‐harvest recruitment patterns were not spatially random; Pinus seedlings initially established in natural grass openings and then proceeded to fill‐in stump patches created by harvesting. Knowledge of spatial pattern should be explicitly incorporated into restoration activities in these forests.     

大兴安岭次生林区优势种落叶松分布格局及竞争作用

竞争是形成特定群落结构、构成分布格局的基本驱动力之一,树种的空间分布和大小并不是相互独立的,而是广泛受到竞争过程影响。在大兴安岭设置一块具有代表性的天然次生林恢复样地,采用点格局分析中的最近邻体距离G（r）函数、双关联函数g（r）和基于个体胸径标记的点格局分析方法同时结合4种点格局零模型（完全随机模型、均值托马斯模型、随机标签模型、先决条件零模型）研究分析大兴安岭次生林区优势种落叶松不同生活史阶段的分布格局、关联性及竞争作用。研究结果显示：落叶松不同发育阶段分布格局不同、同一发育阶段不同研究尺度分布格局也不同;落叶松种群内存在潜在的竞争、且竞争作用与龄级和研究尺度有关;落叶松幼龄个体在小尺度（r<5m）下的聚集分布是由扩散限制作用和竞争作用所导致;落叶松种群内成龄与幼龄个体间有较强的正关联性、成龄个体对幼龄个体在小尺度下有明显的庇护作用;标记点格局分析方法在检测植物群落中的竞争作用时敏感性较高。